Bulletin of the California Lichen Society 2006 13-2

Bulletin
of the
California Lichen Society

Volume 13

No. 2

Winter 2006


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discussions through />Volume 13 (2) of the Bulletin was issued 20 December 2006.

Front cover: Ileodictyon sp. (see article on p. 35). Photography by Darrell Wright.


Bulletin of the California Lichen Society
VOLUME 13

NO. 2

WINTER 2006

Cladonia firma in San Luis Obispo County, California
Kerry Knudsen
The Herbarium, Department of Botany & Plant Sciences
University of California, Riverside, CA 92591-0124


James C. Lendemer
Department of Botany, The Academy of Natural Sciences of Philadelphia
1900 Benjamin Franklin Parkway., Philadelphia, PA, 19103


ABSTRACT
The populations in California are verified as
Cladonia firma sensu stricto.
KEYWORDS
Cladonia firma, Los Osos, Montana de Oro
State Park, San Luis Obispo County, lichens of
California.
Cladonia firma (Nyl.) Nyl. Bot. Z., 1861: 352,
1861. Type: Portugal: Algarve, marim in glareosis

maritimis, elevation about 5 m C.N. Tavares:
Lichenes Lusitaniae selecti exsicatti No. 39 (H!
neotype)
Basionym: Cladonia alcicornis var. firma Nyl.,
Syn. Lich., 1: 191, 1858.
Synonyms: Cladonia foliacea var. firma (Nyl.)
Vain.; Cladonia nylanderi Cout.
INTRODUCTION
Botanist Jeanne Larson first discovered an
unusual Cladonia species with large green cleft
squamules in a vacant lot next to her parent’s home
in 1973 in Los Osos in San Luis Obispo County,
California. Charis Bratt collected specimens in Los
Osos in the 1980s and brought them to the attention
of Cladonia specialists Samuel Hammer and Teuvo
Ahti. Ahti considered the specimens to be C. firma,
previously known only as an old-world species, but

Hammer considered the Los Osos populations to
represent C. firma in a broad sense and felt they
could belong to a separate, though superficially
similar taxon (Hammer, pers. comm.). Despite this
divergence of opinion the Los Osos populations were
published as C. firma (Hammer 1991, 1993) with a
later cautionary note that the identification was not
taxonomically clear because of their divergent
opinions (Ahti and Hammer 2002).
When Cladonia firma was first collected it was
locally abundant in the Los Osos area. Recently
housing developments have spread through the

Baywood-Los Osos area, severely reducing local
habitat and extirpating populations. Existing
populations are separated and several are in decline.
Concern for the conservation of the dwindling
populations in the Los Osos area, spear-headed by
conservationist David Magney and California State
Park ecologists Lisa Andreano and Mike Walgren,
stimulated the authors to undertake a taxonomic
study of the remaining populations to decide whether
they were Cladonia firma in the strict sense or a
species new to science. In this paper, we report the
results of our study. Our observations are based on
field studies conducted in 2006 as well as specimens
collected specifically to represent a full suite of
morphological variation.

29


BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 13 (2), 2006
coating eventually thins or disappears and the cortex
TAXONOMIC AND ECOLOGICAL DESCRIPTION OF
turns dark brown. This can appear in fresh specimens
LOS OSOS SPECIMENS
to have a bluish tint to some people.
The thallus is squamulose and the squamules are
The upper surface of the squamules is a green to
persistent forming small clumps, 2-25 cm in
olive, sometimes becoming brown. It is glaucescent
diameter, often sterile and without podetia when

because of a syncortex in sensu Knudsen (Knudsen in
young. It is conspicuous when dry because the large
prep.), an upper and uneven gelatinous coating up to
squamules roll inward, are upright and densely
100 μm thick, punctuated with pits and valleys where
packed together, exposing white or brown, esorediate
undersides. The primary squamules are
the largest in California, up to 25 mm
long and 10 mm wide, deeply cleft and
digitate
with
often
secondary
a
crenulation. They are up to 250 μm
thick. The crenulations of squamules
elongate into digitate straps at the end
of which new squamules form (Figures
1a and 1b). It is this process of
elongation that gives the species its
complex form (Figure 2). In
undisturbed sites, Cladonia firma forms
contiguous populations. In mildly
disturbed sites, C. firma readily
fragments, eventually forming new
thalli that are tangled, attenuated
structures of interconnected squamules,
stalked pycnidia, and podetia with
secondary squamules. This ability to
easily regenerate, even if turned

completely upside down, is welladapted to the sandy maritime sites C.
b
firma favors where animals, winds or
rainstorms may fragment, flip, or
partially bury individuals (Figure 2).
The thallus does not usually grow
directly on the sand in the Los Osos and
Montana de Oro populations, but
actually favors openings in the maritime
dune scrub or openings formed by the
death of maritime chaparral where the
sand is covered with a thick layer of
detritus and there is abundant rabbit
dung. It also grows on mosses. These
sites are generally level or gently
inclined.
The lower surfaces of the
squamules are corticate with periclinal
prosoplectenchyma and covered with a
thick white fibrous coat of fine hyphae.
In older squamules, this coat can
Figure 1. Typical developments of thalli. Photography by James C.
blacken, probably due to interaction
Lendemer.
with soil or bacteria. Usually the fibrous

30


Cladonia firma

the gelatinous layer is often as thin as 5 μm. This
variation of thickness gives the surface a bumpy
texture which is probably functional because water
accumulates between the thick bumps in lower areas
on the squamule surface and can easily be absorbed
where the gelatinous layer is thin. The eucortex in
sensu Knudsen is formed of mostly anticlinal
prosoplectenchyma and is 30-50 μm thick beneath
the upper syncortex.
The podetia usually begin from the center of the
primary squamules (Figure 1b), arising to a height of
up to 15 mm, sometimes branching, but narrow,
usually 1 mm in diameter. Several podetia can arise
from one squamule. The podetial surface is corticated
and covered with bumps which are nascent
squamules but can develop into new podetia if
flipped over. The podetium is cup-bearing, the cup
usually abruptly flaring out. The cups are usually
shallow, 2-3 mm in diameter, and often one to three
podetia arise from the center to form a second or
third tier, resembling C. cervicornis (Ach.) Flotow.
Sometimes secondary squamules develop around the
rim of the cup or apothecia or pycnidia. It should be
noted that these podetial characters are common in
the C. cervicornis group and are pleisomorphic and

Figure 2. Thallus of Cladonia firma (Nyl.) Nyl.
Photography by James Lendemer.

not unique to C. firma. But the podetium of C. firma

thickens and readily elaborates (see Figure 1).
Most herbarium specimens from California lack
podetia. Actually, this absence is due to collection
bias. In Los Osos, specimens with podetia were
collected separately and identified and reported as C.
cervicornis. Actually the two species can be easily
separated: the squamules of C. firma are distinctly
larger, C. firma contains atranorin which C.
cervicornis s. str. lacks, and C. cervicornis (which is
rare in Los Osos) has larger podetia.
The apothecia are brown and usually develop
sessile or stalked on the rim of cups. The ascospores
are hyaline, simple, and 14-17 x 2-4 µm.
The pycnidia are brown, urn-shaped, sessile or
stalked, arising on the edge of cups, on the sides of
podetia, and from upper surface of primary
squamules. The conidia are sickle-shaped, 5-7 x 1
µm.
Fine rhizohyphae, acting as anchors, can occur
on the underside of thalli.
Comparison with the neotype of C. firma as well
as with a selection of European specimens showed no
morphological divergence in the structure of the
thallus, the podetia, or in conidia, or spores. Thus we
consider the Los Osos-Montana de Oro populations
to be C. firma in the strict sense.
Cladonia firma can be easily determined by its
large primary and persistent squamules, the largest in
California (see Figure 2). The key in Lichen Flora of
the Greater Sonoran Area, Vol. 2 (Ahti and Hammer

2002) works well for determining all Cladonia
collected so far in San Luis Obispo County.
CHEMISTRY
European specimens of C. firma contain
atranorin and fumarprotocetraric acid with smaller
amounts of protocetraric acid and confumarprotocetraric acid, which is part of the
fumarprotocetraric acid chemosyndrome (annotation
of high performance liquid chromatography by K.
Huovinen and T. Ahti in 1985 on Follman: Lichens
Exsiccati Selecti A Museo Historiae Naturalis
Casselensi Editi No. 124, H!)
All specimens from Los Osos were tested with
thin layer chromatography (TLC). HPLC tests of
specimens of C. firma from Los Osos were not
performed. All of the specimens reviewed for this
study contained atranorin as an accessory to
fumarprotocetraric acid. They were consistent in the
production of fumarprotocetraric acid throughout the

31


BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 13 (2), 2006
thallus and inconsistent in the production of atranorin
in all parts of their thalli in high enough
concentrations to be detected by spot tests or TLC.
This led to some samples appearing to lack atranorin
when first run through TLC. Adequate thalli samples
are needed for good atranorin results. Spot tests were
P+ orange, K+ yellow or rarely K- or a dingy

yellowish reaction that is not very conclusive.
There are atranorin-rich populations of an
undescribed species in western North America,
reported by Ahti and Hammer (1990) and included in
C. cervicornis (Ahti and Hammer 2002). This species
occurs from Amador and Lake Counties to Torrey
Pines and Point Loma in San Diego County,
California, in small scattered populations. But it is
abundant on Santa Rosa Island, the center of its
distribution. Like C. cervicornis and C. firma it has
tiered corticate podetia. Like C. firma it has atranorin.
It has smaller, simpler squamules than C. firma. Its
podetia tend to be one-or-two tiered and smaller than
C. cervicornis.
DISTRIBUTION
Cladonia firma is abundant locally in Spain and
Portugal with populations scattered in sandy
maritime habitats around the Mediterranean as well
as on the Canary Islands and the Channel Islands of
Great Britain.
There are five known sites of Cladonia firma in
California (see Figure 3). In recent surveys we have
observed only two major populations, one in Los
Osos and one in Montana de Oro. Both contain
thousands of individuals. We observed two other
populations. One was on a vacant lot in Los Osos
were it occurred on detritus under several decorticate
and lichen-covered shrubs. The other was off
Baywood Heights where according to David Magney
(pers. comm.) about 500 individuals persist scattered

across the lower section of a 70-acre parcel. A third
major population was surveyed by David Magney
south of Los Osos Middle School on Pismo Street.
He has estimated a population of several thousand
thalli on five acres. We have not observed it yet.
The populations in Los Osos and Montana de
Oro are the only known occurrence of the species in
North America. We expect more populations to be
found in Montana de Oro State Park and on scattered
parts of undisturbed land in Los Osos. Though it is
possible new populations will eventually be
discovered in other parts of western North America,
it should be kept in mind that coastal habitats like

32

Los Osos with the unique combination of stabilized
dunes with open maritime scrub and particularly
moist maritime conditions may not be as common or
undisturbed as one might hope.
CONSERVATION
The major threat has been realized and most of
the stabilized dune habitat between the Powell
property in Los Osos and Montana de Oro State Park
has been developed. The remaining populations are
the separated remnants of what was probably once a
large and contiguous local distribution.
The Powell property in Los Osos is divided
between Fish and Game and the state parks. There
are several thousand individuals of C. firma scattered

around the area. It is unfenced, across from a school
with several houses in the area. People regularly walk

Figure 3. Map of distribution of Cladonia firma
in North America.
through the area for recreation, sometimes taking
their dogs with them, probably not more than a few
per day at most. But the sand is very fine and deep.
People walking through the area tend to avoid the
thickets of oak and chaparral and follow the openings
in the maritime scrub where C. firma is most likely to
occur. Just normal walking begins to churn the sand
and suck under detritus, bryophytes and C. firma,
burying them. Dogs do more damage. Even in areas


Cladonia firma
at the Powell property that look undisturbed many
thalli are fragmented, lying on the sand.
The conservation of the populations at Powell
can only be achieved through the fencing off of the
area and any recreational walking, if allowed,
restricted to elevated walkways or fenced trails as is
done at the nearby Elfin Forest Preserve. We hope
one day the management of the Powell properties
will be consolidated and the area given preserve
status.
The population of several thousand individuals
in Montana de Oro, on a sandy ridge above the Sand
Spit area, is undisturbed on beds of detritus, forming

thick stands of contiguous individuals. These look
like the best European specimens we have seen.
Though near a well-churned trail that leads to the
beach, there were no signs of human or animal
disturbance. The Montana de Oro population could
be impacted if a new trail was made through its area
to allow the old trail to be rehabilitated.
Exploring the Los Osos area, it was evident that
much suitable maritime dune scrub habitat is overrun with Veldt grass (Ehrharta calycina Sm.). In
fact, in central California there is supposed to be
more perennial Veldt grass than in its native habitat
in South Africa (Walgren, pers. comm.). This is
probably the single greatest threat to the remaining
populations of C. firma. This invasive grass needs to
be regularly monitored and removed. The maritime
dune scrub at the Baywood Heights site is suffering a
massive invasion of Veldt grass and if it continues we
expect C. firma to eventually disappear from that
property.
CONCLUSION
Though we consider based on morphological and
chemotaxonomic evidence that the Los Osos
populations are Cladonia firma in the strict sense, the
next step in studying this disjunctive occurrence in
North America would ideally be the comparison of
molecular markers of both the fungal and algal
components of the Los Osos populations with
European and Canary Island populations of C. firma.
Though we would not expect to have to re-evaluate
its species concept using different character states

that are not apparent at this time, we do expect such
research to shed light on the fascinating subject of the
lineages of C. firma.
As concerned scientists, to help assure proper
management on public lands of Cladonia firma, we
are sponsoring the species through the listing process

of the California Lichen Society's Conservation
Committee, so that the species may fall under
protections provided by the California Environmental
Quality Act.
SELECTED SPECIMENS
CORSICA. Ajaccio, vägbrant, 1887, Norrlin s.n.
(H) [det. by Nylander]. FRANCE. Dept. Finistère:
Forêt du Cranou, on silicate soil with iron content,
Lambinon 60/F/731 (H); Dept. Vendeè: Ile d’Yeu,
Marais 1754 (H). GREECE. Dalmatien, Poelt s.n. =
Plantae Graecenses 14 (H). PORTGUAL. Algarve,
marim in glareosis maritimis, Tavares s.n. =
Lichenes Lusitaniae selecti exsiccati 39 (H, neotype).
SPAIN. Andalucía: Prov. Huelva. Coastal formation
E of Mazagón near Torre del Oro, Follman s.n. =
Lichenes Exsiccati Selecti A Museo Historiae
Naturalis Casselensi Editi 124 (H). Canary Islands.
Gomera, Vegaipala between Montaña de Yerta and
Tagamiche, Follmann s.n. = Lichenes Exsiccati
Selecti A Museo Historiae Naturalis Casselensi Editi
341 (ASU); Soria. Matalebreras (40 km NEE of
Soria), common on basic soil, Ahti & Burgaz 50776a
(H). USA. California: San Luis Obispo Co.: Los

Osos, state park property (Powell 1), east of
Bayshore Drive, 35° 19' 37"N, 120° 49' 13"W,
locally common on Baywood fine sand in opening of
chaparral, elev. 27 m, Knudsen 5013 & Lendemer
(UCR, hb. Lendemer), Knudsen 4547.1 & Andreano
(UCR); sand dunes, Bratt s.n. = Cladoniaceae
Exsiccatae Americanae 43 (ASU, NY); ridge N of
Baywood, SE of Morro Bay, Bratt 7273 (ASU,
SBBG); s/e corner of South Bay & Nipomo Street,
35° 18' 01"N, 120° 49' 27"W, elev. 36 m, Knudsen
2776 (UCR); Cordoniz property, east of Bayview
Heights & Calle Cordoniz, 35° 18' 19"N, 120° 49'
54"W, elev. 78 m, Knudsen 7352 & Andreano UCR);
Montana de Oro State Park, ridge of stabilized dunes,
35° 17' 58"N, 120° 52' 08"W, elev. 75 m, Knudsen
7261 (H, PH, UCR); Montana de Oro State Park,
Riefner 87-39 (UCR).
ACKNOWLEDGMENTS
We thank Trevor Goward and Eric Petersen for
reviewing this paper. We thank S. Hammer and T.
Ahti for their comments and help, the California State
Parks for access to their properties, Lisa Andreano
and Mike Walgren for aiding us in the field as well as
giving us shelter, David Magney for stimulating our
interest in the first place and encouraging our
investigations and supplying his own unpublished

33

BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 13 (2), 2006
data, as well as J.A. Elix, Charis Bratt, Shirley
Tucker, and the curators of ASU, H, NY, SBBG,
SFSU and UBC for loaning specimens.
LITERATURE CITED
Ahti, T., and Hammer, S. 2002. Cladonia. In: Nash,
TH, III, Ryan, BD, Gries, C, Bungartz, F. (eds.):
Lichen Flora of the Greater Sonoran Desert
Region. I. Lichens Unlimited, Arizona State
University, Tempe, Arizona, pp. 131-158.
Hammer, S., and Ahti, T. 1990. New and interesting
species of Cladonia from California. Mycotaxon
37: 335-348.

Hammer, S. 1991. A preliminary synopsis of the
species of Cladonia in California and adjacent
Oregon. Mycotaxon 40(1): 169-197.
Hammer, S. 1993. A revision of Cladonia section
Perviae in the western United States. Bryologist
96(3): 299-309.
Knudsen, K. in prep. Acarospora In: Nash III, T.H.,
Ryan, B.D., Diederich, P., Gries, C., Bungartz,
F. (eds.): Lichen Flora of the Greater Sonoran
Desert Region, Vol. 3. Lichens Unlimited,
Arizona State University, Tempe, Arizona.

Crustose lichens at Torrey Pines State Reserve: Buellia maritima (A. Massal.) Bagl. (left) and Caloplaca luteominia
(Tuck.) Zahlbr. var luteominia (right). Photography by Rolf Muertter.

34

An Odd Non-Lichen Fungus With an Echo of The Lichens
Darrell Wright
35 Kempton St., Greytown 7512
New Zealand

After dinner at East Taratahi, Carterton, lower
North Island, Ashley Toms brought in the object in
Figure 1 (front cover), which he had found on the
floor of a 10 acre remnant of podocarp forest on the
property. After working out that it was some sort of
fungus, I noted similarities to the lichens Cladia and
Ramalina. The three dimensional net is reminiscent
of Cladia retipora (Figure 2) of New Zealand and
Australia, and the perforations remind one of other
Cladia species and of Ramalina section Fistulariella,
e.g., R. dilacerata (Figure 3) of North America. The
hollow, tubular, more or less cladoniform branches
also suggest the lichens.
A search of the internet turned up some very
good on-line guides to the non-lichen fungi of New
Zealand (there seem to be none of these for the whole
of North America, and one author claimed there is
not even a checklist of North American non-lichen
fungi!). Several of these guides showed the fungus to
be Ileodictyon (intestine and net), Basket Fungus,
perhaps I. gracile because of the expanded nodes
rather than the more common I. cibarium.
Ileodictyon, formerly placed in Clathrus, has a nearly

worldwide distribution but does not seem to be in
North America. I wondered if there might be some
ancestral connection between it and Cladia apart
from the fact that they are both fungi.
Ileodictyon is a member of the Phallaceae, the
stinkhorn fungi, basidiomycetes which are unusual in
that the spores are dispersed not by air currents or
water but by insects which are attracted to the carrion
odors which the fungi produce. What you see in
figures 1, 4, and 5 is called a receptacle or popularly
an “egg” (Maori referred to the “eggs” as tutae
kehua, the feces of ghosts, or tutae whetu, the feces
of stars!). The “eggs” develop in what looks like a
puffball. At maturity the puffball bursts releasing
perhaps 10 “eggs”, which expand into these geodesic
dome type cages (I found comparisons to
Buckminster Fuller balls and to soccer balls with the
panels knocked out). The spores are produced in a
foul smelling gelatinous mass (gleba) on the insides
of the hollow tubes, called “arms,” of which the cage

is formed. I saw no mention of the perforations
(Figures 4 and 5).
Several writers theorized that the fungus would
be dispersed when the “eggs” were blown about (they
are not attached to the substrate). Wind might help to
move the “eggs” over short distances on the forest
floor in tumbleweed fashion but would not account
for the release of the spores which are fixed in the
sticky mass inside the cage members, that is, the

perforations are not for the spores to merely fall
through. I think they must be portals for the insects
which enter the tubular arms of the cage to feed on
the spores and end up getting them stuck to their
bodies. I found an insect larva within one of the arms,
suggesting that insect mothers lay their eggs within
the arms where the eggs are sheltered and there is
food ready and waiting for the emerging larvae.
But has this fungus anything to say about the
lichens? Probably not. First, Ileodictyon is a
basidiomycete while Cladia and Ramalina are
ascomycetes, belonging to that group of fungi, very
different from the basidiomycetes, of which most
lichens are members. Second, the analogous
structures have different ontogenies. In Ileodictyon,
the baskets develop by linking up (anastomosing) of
the arms (Cunningham 1931; some related Clathrus
species have arms which join only at the apex of the
receptacle or not at all), while Hammer (2000) has
shown that the perforations in Cladia retipora
develop from creases in the branches, which
correspond roughly to the “arms” of Ileodictyon.
Third, the analogous structures have different
functions. In Ileodictyon they are bound up with
reproduction and dispersal (Cunningham 1931). As
far as is known there is no connection between
dispersal and the perforations of Cladia and
Ramalina. The tiny perforations of Cladia spp. other
than C. retipora and of Ramalina section Fistulariella
may help to hydrate the thallus by admitting

atmospheric moisture to the interior in dry weather.
Many Hypogymnia and Cladonia species have
perforations which could perform a similar function,
and Menegazzia is famous for them. This looks like

35


BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 13 (2), 2006

1. [On front cover] Ileodictyon cf. gracile Berk., D.M. Wright 7808, East Taratahi, Carterton, New Zealand.
Receptacle or “egg”.
2. Cladia retipora (Labill.) Nyl., D.M. Wright 7646, Abel Tasman National Park, New Zealand.
3. Ramalina dilacerata (Hoffm.) Hoffm. (Fistulariella minuscula [Nyl.] Bowler & Rundel), D.M. Wright 7099,
Elk River, Humboldt Co., California.
4. Ileodictyon 7808, arms with perforations at nodes.
5. As for 4, close-up of perforations, some of which contain rainwater.
an instance of convergent evolution, of unrelated or
distantly related organisms which have independently
developed similar morphologies, in this case by
different routes and for different purposes.
However, if, during its further evolution,
Ileodictyon were to meet a nice compatible alga...
LITERATURE CITED
Cunningham, G.N. 1931. The Gasteromycetes of
Australasia. XI. The Phallales, part II.

36

Proceedings of the Linnean Society of New

South Wales 56(3): 182-200. Available at
/>/data.asp, accessed 22 July 2006.
Hammer, S. 2000. Meristem growth dynamics and
branching patterns in the Cladoniaceae.
American Journal of Botany 87: 33-47.
Available
at
http//www.amjbot.org/cgi
/content/full /87/1/33, accessed 22 July 2006.


Notes on the Lichen Flora of California #3
Kerry Knudsen
The Herbarium, Department of Botany & Plant Sciences,
University of California, Riverside, CA 92521-0124.
ABSTRACT
Terricolous lichen habitat and terricolous
species are discussed in California.
KEYWORDS
Acarospora
schleicheri,
Acarospora
terricola, Casper Wilderness Park, Cladonia
nashii, lichens of California, Santa Rosa
Plateau, terricolous lichens.
Working in the field in southern California, I am
impressed with how extensively terricolous lichen
habitats have been destroyed. Before Thanksgiving I
visited the Casper Wilderness, an Orange County
park, in the coastal foothills of the Santa Ana

Mountains to take a look at the local lichen flora with
Senior Ranger Donna Krucki and Ranger Joy Barnes,
who is moving to Washington in December.
In the openings of the coastal sage scrub and
chaparral, non-native grasses, dried out by the long
hot summer, were dominant. Ecologically these
ruderal grasses are especially dangerous because they
add a fuel load to an area that is susceptible to fires.
These grasses, and the weeds associated with them
like Eurasian mustards and star thistles in moister
areas, have taken over a habitat that once supported a
flora of native annuals and native bulbs as well as
terricolous lichens and native grasses.
Donna, a ranger at the park for twenty years, led
Joy and me to an area where she had seen lichens on
soil. The site was sad. The area was clay. On the
edges of eroded runnels, Cladonia nashii Ahti was
reduced to depauperate squamules and scattered
stunted podetia hard to recognize from other species.
Cladonia nashii is apparently endemic to California
and is a member of the Cladonia humilis group as is
the closely related Cladonia hammeri Ahti. Cladonia
nashii, despite the description in the Sonoran flora
that it has farinose soredia (based on a handful of
specimens) (Ahti and Hammer 2002), eventually
develops consoredia on its short expanded dentate
cups that form granulose structures which are
ecorticate piles of stringy hyphae and algae in clear
globes of gelatinized hyphae. When the cups are

disturbed the sorediate gobs are knocked loose, but
more often in rain when the cups fill with water, the
granulose soredia float loose and eventually spill or
are splashed on to the soil spreading the species.
Cladonia hammeri is almost indistinguishable
because it has granulose soredia too but these
granules tend to be corticate. Cladonia nashii
contains atranorin which can usually be seen if one
allows bubbles of K to form on the podetia which are
a rich yellow that does not turn a murky red. Rarely
is TLC needed. Cladonia hammeri does not contain
atranorin.
Anyway, seeing these poor Cladonia clinging to
the last tiny niches and barely thriving, I lost hope in
Donna showing me any interesting soil lichens.
Instead I collected lichens off a pile of sandstone
rocks deposited by erosion, showing each one to the
rangers and telling stories and answering questions.
But Donna, aware I was disappointed, urged us
on to see another site. We bushwhacked and I was
thinking I needed to get away from my microscope
and computer more as I sweated in the unusually hot
November afternoon and chaparral branches stabbed
me in the legs. Eventually we found the trail Donna
was looking for and climbed up a long ridge. On its
long lanky peak were barren white patches where
eroded sandstone formed thin and nutritionally poor
soil over bedrock with scattered nodules of rusty
iron-rich sandstone. Donna and Joy told me that most
of the rare annuals discovered in the park had been

found on these barren patches which I could see on
other ridges.
I immediately recognized the rare soil lichen I
have been studying, the brown Acarospora terricola
H. Magn. (see the upcoming book on soil crusts by
Bruce McCune and Roger Rosentreter for excellent
pictures). It can look a little like the pruinose Psora
pacifica Timdal that was growing nearby. This was
only the seventh site I have found A. terricola in
California and I know of only three specimens from
other locations, one in southern California near
Banning collected by Wetmore, and one specimen
each from Washington and Nevada. The type

37


BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 13 (2), 2006
specimen was originally collected by Hasse in the
Santa Monica Mountains (W!).
On the white patch on the ridge was the
brownish Lecidea fuscoatra (L.) Ach. But
everywhere was a lichen with black lecideine
apothecia too but with a thin whitish thallus barely
emerging from the sandstone. These turned out to
belong to an apparently undescribed Sarcogyne sp. I
have never seen before though I regularly collect and
study the genus. It may be related to a specimen that
Bruce McCune has found in Oregon forming a
thallus.

Carefully walking forward, suddenly I saw the
yellow Acarospora schleicheri (Ach.) A. Massal. and
got really excited. Based on herbarium collections
and Hasse (1913) it was once common in southern
California. After four years of surveying I have still
not found it in the Santa Monica Mountains. It was
once common according to Hasse in Lake Elsinore
too near where Liz and I live but after years of
collecting in this area I know of only one site in the
Menifee Hills. And I have only found one site in the
Santa Ana Mountains so far. Acarospora schleicheri
is a beautiful granular yellow crust with many dark
discs. I no longer glue collections of it because I am
interested in its dense anastomosing rhizohyphae
which grow underground and slowly spread a
population outward. It is especially sensitive to any
disturbance as is A. terricola. Even one too many
visits by lichenologists can lead to its demise.
As I drove back over the Santa Ana Mountains,
playing over and over a new ragged hard-driving
song by Bob Dylan, I didn’t even hear the music as I
dreamed of what southern California must have
looked like before the Spanish came with cattle and
the first weeds. But such thoughts have a hard sad
edge that cuts deep and an awareness of an
irrevocably changing natural environment can lead to
cynical feelings of hopelessness and a tendency to
project into nature an ugliness that can blind us to
those high white ridges of hope that rise above the
grass-coughed slopes.

A week later, as December began, I was out
hiking on Santa Rosa Plateau more out of a resolution
to get more exercise than to collect though I had a
permit and my collecting backpack was heavy with
hammers and chisels, shears and saws. The plateau,
famous for its vernal ponds, was once the Santa Rosa
Ranch. Native grasslands were reduced to ruderal
grasslands by decades and decades of grazing.
Listening to my friend, the resource manager Carole

38

Bell, talk about the plateau I was deeply impressed at
what a long hard struggle it is to restore the native
grasslands and to prevent weeds from building up
fuel loads that could destroy the oaks and chaparral in
a catastrophic fire.
Looping back toward my car across the Mesa de
Colorado, I went off trail to examine the lichens on
the basalt outcrops that stick out of a heavy clay soil
which lies thinly over a basalt bedrock that supports
vernal ponds. Poking through the grass, I was
surprised to see in a reddish opening a small thriving
population of yellow Acarospora schleicheri growing
with a sterile imbricate Placidium sp. and
Trapeliopsis glaucopholis (Nyl. ex Hasse) Printzen &
McCune. This whole mesa had been heavily grazed
for almost a century and if the propagules of these
lichens hadn’t blown in, then remnants had managed
to survive around the edges of the low protruding

basalt outcrops.
The soil lichens were coming back.
Next day I saw Carole Bell and while we were
driving in her distinctive purple truck over to see a
prescribed burn site I asked her when they had
stopped grazing cattle on Mesa de Colorado. She said
1986. I told her about the recovering terricolous
lichen community. She became concerned and said
they had burned there maybe in the early nineties to
knock back the non-native grasses. Would a
prescribed burn hurt the lichens? Of course it would.
But next time, if it becomes necessary to do a
prescribed burn, she’ll make sure there are cleared
areas around the soil communities. They will survive
and thrive.
There’s hope.
ACKNOWLEDGMENTS
Special thanks to Amanda Heinrich for
reviewing this paper. Special thanks to Joy Barnes,
Donna Krucki and Carole Bell, women out there
making a difference.
LITERATURE CITED
Ahti, T., and Hammer, S. 2002. Cladonia. In: Nash,
TH, III, B.D. Ryan, C. Gries, F. Bungartz (eds.):
Lichen Flora of the Greater Sonoran Desert
Region. I. Lichens Unlimited, Arizona State
University, Tempe, Arizona, pp. 131-158.
Hasse, H E 1913: The lichen flora of southern
California. Contributions from the United States
National Herbarium 17: 1-132.

Mutualism in Lichen Symbiosis
Janet Doell
1200 Brickyard Way #302
Pt. Richmond, CA 94801

Ever since Simon Schwendener determined that
lichens were a combination of a fungus and an alga in
1869, lichenologists have had differing opinions
about the type of symbiosis lichens represented. For
many years the prevailing thought was that the
fungus was parasitizing the alga although not to the
point of killing it; in other words, controlled
parasitism. But that determination was not
unanimous. There were always some who felt that for
a lichen symbiosis to have been so successful and
continuous over time it must have been mutually
beneficial – a sample of mutualism. So even though
Ernie Brodo, in his big volume “Lichens of North
America,” stuck with controlled parasitism, Tom
Nash saw the lichen symbiosis as “a classical case of
mutualism,” as did Rosemary Honneger in 1991, and
Dr. Larry St. Clair of Brigham Young University in
his lecture at San Francisco State on May 10, 2006,
entitled “The Case for Mutualism” and prepared in
collaboration with Kathleen Knight, also from
Brigham Young.
After an introduction which answered the
questions “what are lichens, where are they found

(everywhere) and what do they grow on
(everything),” Dr. St. Claire embarked on his main
point by showing a very good illustration (photo by
Ahmadjian) of an alga being clasped and held by
fungal hyphae, at which point he explained that if the
alga was not the right one in the eyes of the fungus,
the fungus would kill it and look for another one. In
other words, the two partners have to be compatible
for a lichen thallus to evolve from the combination.
As to when lichens first appeared on earth, the
fossil record is poor. But based on the fact that both
algae and fungi had to be present for this symbiosis
to exist, lichens probably appeared in the Silurian
period, about 440 million years ago. The most
primitive lichens are leprose ones. These lichens have
no real thallus but consist of granules made up of a
few fungal hyphae and a few algae. Fruticose lichens
with their more complex thalli followed, as did the
foliose and crustose genera, each introduced in this

lecture by illustrations of their distinctive structures.
For instance, the lower surface of crustose lichens
does not have a lower cortex and the fungal hyphae
can penetrate the rock substrate to a depth of 10
millimeters.
Algae reproduce only by cell division within the
lichen thallus; and the fungus also reproduces largely
by vegetative means. Vegetative reproduction in
lichens is aided by soredia, small granules of a few
fungal hyphae and a few algae which are easily

spread by wind or animals or rain, and by isidia,
small finger like projections from the thallus which
are designed to break off and spread the lichen to
new areas, carrying a bit of cortex along as well as
the fungus and the alga.
The fungus does reproduce sexually in many
lichens, and Dr. St. Claire showed diagrams of the
apothecia where the spores develop. When the spores
are mature and become scattered they must find an
alga to capture before they can develop into a lichen.
Interestingly enough, the algal genus Trebouxia is
rarely if ever found living independently in nature,
even though it is the alga most commonly found as a
photobiont in lichens.
Getting down to the nitty gritty of symbiosis,
which partner benefits the most in this arrangement
and which one provides the most? Who provides
what? Is it mutual?
What does the alga contribute?
1. The alga, as photobiont and thus the one
who does the photosynthesizing, produces fixed
carbon, and transfers 90% of its carbon
production to the fungus, providing its partner
with energy and the building material it needs
for growth and repair. Fungi are made of chitin,
the production of which requires a lot of carbon.
The algae produce more fixed carbon overall
when in this symbiotic relationship than when
alone.

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BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 13 (2), 2006
2. In addition, about 5% of lichens have
cyanobacteria for their photobiont instead of
algae, and they are able to fix nitrogen as well as
carbon and pass that on to the fungus to help
meet basic metabolic needs. There are also some
lichens which have basically green algae for
their primary photobiont, but also have
structures called cephalodia, small outgrowths
from the thallus, in which cyanobacteria are
located.
In short, the alga contributes carbon, and in some
cases, nitrogen.
What does the fungus contribute?
1. Nutrient transport. When the lichen thallus
is saturated with water, as it is on a regular basis
under ideal conditions, it would be very difficult
for CO2 to be available for the algae to fix it and
thus make it available to the fungus. To
circumvent this problem, the fungus forms
conduits along the inside walls of the fungal
hyphae in the medulla, and strengthens the walls
of this canal by depositing a layer of
hydrophobic proteins along the outer wall. Water
and nutrients can thus be moved up to the algal
layer from the soil layer while keeping the
medulla dry enough for CO2 to be available to

the algae.
2. Water storage. The fungal hyphae will suck
calcium out of the rock substrate and turn it into
calcium oxalate crystals. Some of the water in
the thallus is bound to the calcium oxalate, and
thus stored for future use. For instance, the
fungus can store some of the water deposited by
the morning dew, and then release it later in the
day so that the alga can continue to
photosynthesize when the lichen is dry. The
fungal partner has evolved so as to keep the
photobiont with enough dampness to continue
working. Green algae can photosynthesize with
water vapor. Cyanobacteria need liquid water in
order to photosynthesize.
3. Gas exchange. Gas exchange of any kind is
difficult when there is too much water in the
thallus. In addition to helping with CO2
availability as seen in paragraph one, the fungus
also has structures on lichen surfaces which
allow gas exchange, in the form of small
openings in either the upper or lower cortex of

40

many lichens. These openings are called
cyphellae and pseudocyphellae and vary in
shape.
4. Light regulation. Calcium oxalate is also
deposited as a dust on the surface of the lichen,

acting as a kind of sun screen to protect the algae
from too much light. Calcium oxalate crystals
also affect light by reflecting it back and
transforming ultraviolet light into enough white
light to allow the algae to photosynthesize more
hours of the day. Pigments in the fungus also
filter and regulate the amount of light coming
into the thallus.
5. Distribution of elements. Phosphorous,
chlorine, sulfur and other nutrients are delivered
to the algal layer via the canals discussed earlier.
6. Herbivory. Invertebrates like to graze on the
lichen algae. The fungus and the alga together
produce secondary chemicals which help fight
these invaders.
7. Pollution protection. The lichen fungi can
neutralize polluting elements that they absorb
and store them. Sulfur, arsenic and lead are
sequestered in crystal structures to keep them
from affecting the algae.
8. Reproduction aids. Fungal and algal
components cooperate in forming asexual
diaspores like isidia and soredia, which are
designed to keep the alga together with the
fungus in vegetative reproduction. In some
perithecial lichens there are algal cells near the
perithecia and spores are sometimes scattered
with algal cells on their surface. In one such
genus algae are found in the hymenial layer and
fungal spores can be discharged with algae

attached. In other words, algae can get to places
in this symbiotic arrangement that they would
never get to otherwise.
The fungus seems to know that it has a good
thing going, and it treats its algae well. In return, the
alga gives up 90% of its product to the fungus. As
Dr. St. Clair put it, “The rent is high, but the returns
are incredible for the alga.”
Even though the alga provides all the food, if the
fungus is aiding it in eight different ways to lead a
comfortable life, the alga resembles a prisoner less
than it does a musician in the 18th century who was
provided luxurious living conditions in return for


Mutualism in Lichen Symbiosis
providing the nobleman who employed with his daily
dose of original live music. This certainly does sound
like mutualism, unless the alga harbors dreams of
freedom, which seems unlikely.
A short question period followed the lecture.
Subjects discussed included procedure used in
separating the alga from the fungus in lichens;
nomenclature; other symbiotic relationships; the lack

of any exchange of genetic material between the alga
and the fungus; the importance of the environment in
lichen success; and the fact that cooperation has been
found to be more common than competition in the
wider natural world.

Thank you, Dr. St. Clair, for taking the time to
present such an interesting and informative lecture.

A Note from the Conservation Committee
Over the last few years, the CALS Conservation Committee has been working to develop and implement a
strategy for ranking and listing sensitive lichen taxa. In doing so, we have been working closely with the California
Department of Fish and Game’s Natural Heritage Program, also known as the California Natural Diversity Database
(CNDDB), and with the California Native Plant Society (CNPS). As you can see in the pages of this Bulletin, our
efforts are beginning to fruit.
California has relatively strong laws for regulating land use in light of rare species. This can bring great
scrutiny to listing efforts. At the core of our process is the sponsorship of taxa. This sponsorship is a gathering of
background information and known locations within California. Any individual may sponsor a lichen. Recognizing
the lack of lichen societies in some states and the greater geographic interests of our own lichen society, we do not
restrict sponsorships to the state of California.
When the committee receives a sponsorship, it is reviewed, then presumably accepted by the committee. The
sponsorship is then publicized by the CALS email group and letters to stake-holders. Publication in the Bulletin as
you see here is not required, but may become commonplace. After publicizing the sponsorship, we generally ask for
comments on technical aspects within 90 days, and general review with opportunity for added surveys over a period
of one year. After this public review, the committee may act to establish a CALS ranking and listing for the lichen.
Our ranking system is built upon Natural Heritage methodology, in use across much of the Americas
(), and functions as a recommendation to the CNDDB. Our listing system is
synonymous with CNPS listings.
For more information on our process, go to . Additional information on CNDDB
and their ranks is available at and on CNPS and their lists at
under the Rare Plant Inventory. If you wish to contribute a sponsorship, then please
coordinate with the committee so as to not duplicate an effort already underway. Contact the committee
chairperson, Eric Peterson,
To date, we have accepted 6 completed sponsorships, and at least as many are in progress. Two of those
sponsorships have completed the one year review and we are happy to announce the first species officially ranked
and listed by CALS. Usnea longissima has received the global rank G5.2, the state rank S4.2, and has been placed

on list 4 (watch). Solorina spongiosa was sponsored for multiple states, receiving the global rank G4G5.3; state
ranks S1.2 (AZ), S1.2 (CA), S1.1 (NV), and S2?.2 (UT); and placed on list 2 (rare in state, but more common
elsewhere) for all four states.
P.S. Precision location data is removed from specimen lists of sensitive species in order to protect them from
poachers and vandals. Yes, that has happened due to publication of data on rare species! The Conservation
Committee maintains these data and can make them available for serious scientific needs.

41


BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 13 (2), 2006

Hypogymnia schizidiata,
Sponsorship for the CALS Conservation Committee
Bruce McCune
Dept. of Botany & Plant Pathology, Cordley 2082,
Oregon State University,
Corvallis OR 97331-2902

EXECUTIVE SUMMARY
Hypogymnia schizidiata (Island Tube Lichen) is
a recently described species that is rare
throughout its limited range, from Cedros and
Guadalupe Islands in Mexico to Santa Rosa
Island and Santa Cruz Island in California. Only
four collections are known from the U.S. The
species should be sought further on the Channel
Islands and populations documented for size,
habitat, and potential threats. At this time, no
imminent threats are known, apart from an

unpopular piece of proposed legislation to
convert Santa Rosa Island to a military
recreation area.
TAXONOMY
Accepted scientific name: Hypogymnia schizidiata
McCune.
Common name: Island Tube Lichen.
Type specimen and location: MEXICO: Baja
California, Cedros Island, south end, top of hill north
of town of Cedros, 28o 8’N, 115o 13’W, 1100 m, on
Juniperus californicus, with Pachycormus, Ambrosia,
and Mammillaria, Marsh 7384, 21 March 1994
(ASU).
Synonyms: None.
DESCRIPTION
Thallus: appressed to suberect, up to 4 (-8) cm in
diam.; texture: cartilaginous; branching: isotomic
dichotomous; budding: absent or rare; lobes: separate
to centrally subcontiguous, 1-3 (-4) mm broad; black
border: not visible from above; profile: even to ±
nodulose; width/height ratio: 1-4; tips and axils:
perforate, upper surface: greenish gray, greenish, or
dark brownish green, often dark mottled, smooth or
becoming strongly rugose; schizidia: formed as
flakes of cortex plus algal layer, developing from

42

either the smooth or rugose areas of the thallus; soredia: sometimes developing on the edges of the schizidia, particularly in rugose areas of the upper
surface, rarely spreading into diffuse laminal soredia;

isidia: absent; lobules: rare; medulla: hollow, ceiling
of cavity brownish to white, floor of cavity brownish
to white; lower surface: black, sparsely perforate;
Apothecia: occasional, substipitate to stipitate, up to
6 mm diam; stipe: urn- to funnel-shaped, hollow;
disc: light to dark brown; ascospores: ellipsoid, (5.5-)
6.5-7 (-8) x (3.5-) 4-5 μm; Pycnidia: common;
conidia: rod-shaped to weakly bifusiform, 6-7.5 x
0.5-0.7 μm; Spot tests: cortex K+ yellow, C-, KC-,
P+ pale yellow, UV-; medulla K-, C-, KC+ orangered, P-; Secondary metabolites: upper cortex with
atranorin and chloroatranorin; medulla with physodic
acid (major), 2’-O-methylphysodic acid (minor), 3hydroxyphysodic acid (accessory, frequency 25%),
unknown C7 (minor UV+ accessory), unknown C8
(minor accessory). Figure 1 (Figure A on back
cover).
Similar species and distinguishing characteristics:
Production of asexual propagules is rather variable,
but in most specimens the cortex tends flake off from
both smooth parts and rugose parts, taking with it the
algal layer. The type is not as heavily schizidiate as
some specimens, but it is fertile and shows the
variation from smooth to rugose lobes. True soredia
can be nonexistent to well developed. The species is
also unusual in its light brownish lobe cavities. In
most specimens it is somewhat variable, having pure
white portions, occasional dark brown portions, but
the majority of the lobe interiors have a loose mesh
of brown hyphae over a white background. This plus
the frequently perforate lobe tips separate the species
from H. imshaugii. The P- medulla is also useful in

separating H. schizidiata from the P+ chemotype of
H. imshaugii. The somewhat darkened lobe cavities
and perforations suggest H. inactiva, another Pspecies, but that species lacks asexual propagules and


Hypogymnia schizidiata Sponsorship

Figure 1 (repeated in color on back cover). Hypogymnia schizidiata McCune. Bratt 3160a from Santa Cruz
Island, showing lobes, perforation in lower surface of lobe tip, and schizidia forming on upper surface of lobe.
Ruler has mm marks. Scale bar in lower right is 1 mm. Photography by B. McCune.
differs in accessory secondary metabolites. Small
pre-sorediate specimens can be recognized by their
brownish cavities, isotomic dichotomous branching,
and perforations in the lobe tips and lower surface.

POPULATION TRENDS
Unknown.
THREATS
History: Unknown.

BIOLOGICAL CHARACTERISTICS
Growth form: foliose lichen.
Reproductive method: typically by schizidia and
soredia; more rarely by ascospores.
Dispersal agents: unknown, but presumably wind,
birds, and arthropods.
Substrate and specificity: on bark and wood of
both hardwoods and conifers, including Juniperus,
Pachycormus, Pinus, Quercus, and Simmondsia.
Habitat and specificity: in woodlands, isolated

groves, and forests.
Pollution sensitivity: unknown.
Ecological function: unknown.
GEOGRAPHY
Global: Offshore islands of Baja California (Cedros,
Guadalupe) and California (Santa Rosa and Santa
Cruz).
Local: In California known only from two collections on Santa Rosa Island, and two collections on
Santa Cruz Island.

Future: Although widely opposed, legislation has
recently (2006) been proposed in Congress to use
Santa Rosa Island for military recreation. It is unclear what this would mean in terms of protection for
the island ecosystems, but presumably it would be
less protected under management for recreation than
for management under the National Park Service
mandate.
Cedros Island apparently has
the largest population of this
species. Cedros Island is well
known for a high level of
endemism. Cedros is remote,
isolated, and hard to reach. It is 24 miles long and
has about 2700 inhabitants. Apparently many of the
inhabitants are unemployed because of changes in the
fishing industry. The main industries are fishing and
a large salt works. Ownership of the sites where the
species occurs is unknown. Cedros Islands is part of
a proposal to create a Biosphere Reserve. This will
include all the 13 islands of the Pacific coast of Baja

43


BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 13 (2), 2006
California.
Presumably this Biosphere Reserve
would provide some measure of long-term protection
for Hypogymnia schizidiata, but at this point it cannot
be considered secure.

Santa Barbara Botanical Garden
ATTN: Charis Bratt
1212 Mission Canyon Road
Santa Barbara, CA 93105

PROTECTION
In 1986 the Federal government purchased Santa
Rosa Island and designated it part of Channel Islands
National Park. In time the island was to become part
of our national parks system, but this has not yet been
accomplished. See threats above.

California Dept Fish and Game
Attn: Roxanne Bittman, CNDDB lead botanist
Wildlife & Habitat Analysis Branch
1807 13th Street Suite 2002
Sacramento, CA 95814

CONSERVATION STATUS SUMMARY

Efforts should be made to document size and status
of populations on Santa Rosa Island and to search for
it on similar Channel Islands. At present, no one has
made a focused search for the species, the existing
records merely being incidental collections before the
species was described.
SPECIFIC CONSERVATION RECOMMENDATIONS
Recommended Global Rarity Rank: G2
Recommended Global Threat Rank: .3
Recommended Local Rarity Rank: S1
Recommended Local Threat Rank: .3
Recommended List: 1B
Recommended conservation/management actions:
Document size and status of populations on Santa
Rosa Island and search for it on similar Channel
Islands.
RELEVANT EXPERTS AND KNOWLEDGABLE LOCAL
BOTANISTS
Bruce McCune, Dept. of Botany & Plant Pathology,
Cordley 2082, Oregon State University, Corvallis OR
97331-2902 USA.
Kerry Knudsen, Herbarium, Dept. of Botany & Plant
Sciences, University of California, Riverside, CA
92591-0124 USA.
STAKEHOLDERS FOR NOTIFICATION OF COMMENT
PERIOD
Channel Islands National Park
1901 Spinnaker Dr.
Ventura, CA 93001

44

California Native Plant Society
Attn: Kristi Lazar, Rare Plant Botanist
2707 K Street, Suite 1
Sacramento, CA 95816-5113
LOCATION/SPECIMEN LIST
Specimens other than the type specimen: MEXICO:
Baja California, Cedros Island, Marsh 7383, 7388
(ASU), 28o 8’N, 115o 13’W, 1000 m, on
Pachycormus, Nash 34483; on Simmondsia, 34497
(ASU); Cerro Redondo, N-facing slope, 800 m,
Marsh 7409 (ASU); N end of island, 28o 22’N, 115o
15’W, 400 m, on Pinus muricata, Nash 34257
(ASU); 442-488 m, Pinus muricata forest,
Beauchamp 62444 (COLO); 28o 8’N, 115o 14’W, on
Juniperus, Nash 34501 (ASU); 28o 8’N, 115o 13’W,
1050 m, on Juniperus, Moran 10647a (ASU, COLO);
Guadalupe Island, 1200 m, Cupressus forest, Tretiach
31618b,c (TSB). USA: California, Santa Barbara
County, Santa Rosa Island, 33o 56’N, 120o 7’W, 480
m, Ryan 31429 (ASU); pine-oak-chaparral mixture,
33o 59’N, 120o 4’W, 260 m, on Pinus remorata, Nash
33051a (ASU); Santa Cruz Island, Bratt 1469. 3160a
(SBBG).
LITERATURE CITED
McCune, B. 2002, Hypogymnia. Pages 228-238 in
Nash, T. H., III, B. D. Ryan, C. Gries, & F.
Bungartz. Lichen Flora of the Greater Sonoran
Desert Region. Vol. 1. Lichens Unlimited,

Tempe, Arizona.
--Editorial Note: Portions of this paper are repeated
from McCune (2002) with permission from the book’s
lead author and publisher, Tom Nash, III.


Sulcaria badia,
Sponsorship for the CALS Conservation Committee
Tom Carlberg
1959 Peninsula Drive
Arcata, CA 95521


David Toren
50 Gardenside #2
San Francisco, CA 94131

EXECUTIVE SUMMARY
Sulcaria badia is endemic to the Pacific
Northwest with an historical distribution from the
Olympic Peninsula in Washington in the north to
the general area of Laytonville, California in the
south, a range of approximately 585 miles. The
contemporary distribution is from Corvallis,
Oregon to Lake Pillsbury, CA, a distance of
approximately 360 miles. In California it is found
in well-lighted Quercus garryana grassland
communities, and in mature Douglas-fir forests
containing a black oak component. Post-2002
discoveries have greatly increased the number

of localities and occurrences in California,
raising the possibility that this lichen is undercollected or overlooked. It is probably dispersal
limited, since neither sexual nor asexual
reproductive structures are known in the
species. Population trends are unknown, since
no monitoring is taking place, although two
historical occurrences are presumed to be
extirpated. It is a Sensitive plant on four National
Forests in northern California.
TAXONOMY
Accepted scientific name: Sulcaria badia Brodo &
D. Hawksw.
Common name: bay
beard lichen

horsehair

lichen,

grooved

Type specimen: holotype F. Sipe #669, OSC.
Type location: U.S.A., Oregon, Philomath. This
population was extant in 2004 (McCune 2004)
Synonyms: none.

Note: Brodo & Hawksworth (1977) state that the
placement of this species into the genus Sulcaria
should be regarded as tentative until fertile material
can be located. The writing of most contemporary

authors treats the species as though that placement
were permanent.
DESCRIPTION
“Thallus pendent, 20–50 cm long, flaccid; branching
mainly isotomic-dichotomous, angles between the
dichotomies mainly acute and rounded; branches
markedly flattened and twisted, conspicuously
sulcate, 0.25 - 0.4 (1.0) mm diameter at the base, with
short, slender almost perpendicular lateral branches;
dull chestnut-brown to almost badious or yellowishbrown in parts. True lateral spinules, isidia and
soralia absent. Pseudocyphellae conspicuous, white,
linear, extremely long, most developing into deep
furrows (Figure 1; Figure B on back cover).
Apothecia and pycnidia unknown. Acetone extract
K+ yellow, C-, KC+ yellow, P+ yellow or brownish”
(from Brodo & Hawksworth 1977). From a distance
of ten feet, the color is the most distinctive character,
although it can be as brown as many Bryoria, and
when pale can still be confused with Bryoria
capillaris.
Similar species and distinguishing characteristics:
At a magnification of 10-14x, well-developed
specimens of Sulcaria badia are easily distinguished
from other Alectorioid lichens by the deep grooves
(pseudocyphellae) that spiral around the main
branches. The color of pale forms from shaded
locales can be very close to Bryoria capillaris, while
thalli from exposed locations can be as dark brown as

45

BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 13 (2), 2006
many other species of Bryoria. Several species of
Bryoria also have pseudocyphellae, but not as deep
or long as found in S.. badia. Bryoria with
pseudocyphellae differ chemically, as well: B.
pseudocapillaris is C+ pink, and B. spiralifera is K+
red. Alectoria sarmentosa can resemble S. badia, but
has a stiffer, more robust cortex, thicker branches (to
2.5mm), and its pseudocyphellae are typically raised,
as opposed to sunken in S. badia. Sulcaria isidiifera
is outwardly similar, but has a different color, and the
pseudocyphellae eventually split open and develop
isidia. It is also distinguished geographically: S.
isidiifera is restricted to San Luis Obispo County,
while S. badia is not known further south than Lake
Pillsbury.
BIOLOGICAL CHARACTERISTICS
Growth form: Fruticose, pendulous, filamentous.
Reproductive method: Primarily
asexual;
via
thallus fragmentation. Apothecia unknown.
Dispersal agents: Birds(?), wind, gravity. Limited

Figure 1 (repeated in color on back cover).
Sulcaria badia Brodo & D. Hawksw. Main
branches (left), showing flattened axil and
spiraling pseudocyphellae, and twisted branches

(right). Photography by T. Carlberg.

46

dispersal ability.
Substrate and specificity: On trees, especially
apple and oak trees (Brodo & Hawksworth 1977). It
is also found on oaks at sites at Lake Pillsbury and on
the Medford BLM District. In northern California on
the Klamath and Trinity Rivers, it has been found as
litterfall in Douglas-fir/black oak forests. Other
substrates include ponderosa pine, Pinus ponderosa,
Douglas-fir, Pseudotsuga menziesii, Oregon ash,
Fraxinus
oregana,
bigleaf
maple,
Acer
macrophyllum, and Rhododendron macrophyllum.
Habitat and specificity: Appears to be a generalist,
but is most abundant in the habitat described by
Brodo & Hawksworth (1977): “on trees, especially
apple and oak trees, in well-lighted Quercus
garryana communities”. The habitats at Laytonville,
Lake Pillsbury (Figure 3) and Medford BLM
localities are consistent with this description,
although the substrates sometimes differ and there is
no summer fog at either of the BLM locations
(Wineteer 2004). The localities proximal to the
Klamath and Trinity Rivers are mid- to late-mature

Douglas-fir forests, sometimes with dense sheltering
canopy covers and either a sparse presence of black
oak (Quercus kelloggii), or none. One Oregon site is
a coastal dune forest.
Pollution sensitivity: Unknown.
Rcological function: Unknown.
GEOGRAPHY
Global: Sulcaria badia is endemic to the Pacific
Northwest, known only from thirteen historic and
contemporary localities in the United States; in
Washington, Oregon and northern California (Figure
2). None of the known localities is further than 85
miles from the ocean. The historic range covers
approximately 605 miles, however the occurrence in
Washington, at the north end of the Olympic
Peninsula, was not relocated by Peterson et al. in
1998, and S. badia has not been found elsewhere in
the vicinity (Hutten 2004) and is possibly extirpated.
The range of extant localities is approximately 360
miles, from Corvallis, Oregon to Lake Pillsbury,
California. One historic occurrence in California is
possibly extirpated (see below), resulting in ten
extant localities for the state. Some localities have
multiple occurrences (see below). The Snider and
Berry Creek localities are the furthest inland. Both
are hot, dry Quercus garryana oak woodlands along
perennial creeks, and both host substantial


Sulcaria badia Sponsorship

populations of S. badia (>¼ mile of creek-side or
>1acre of area) (Wineteer 2004). There are two
specimens in the Arizona State Lichen Herbarium
(ASU) from the Cascade Mountains in Oregon and
California. This would appear to be a significant
disjunct, but see below.
Local (California): There are seven historic and
contemporary general localities in California where
S. badia is known to occur (Figure 2). Some localities
are composed of more than one site where Sulcaria
badia was found, e.g. the Lake Pillsbury area has
nine sites (Toren & Nilles 2003), Hawkins Creek and
Campbell Ridge each have two closely-spaced
occurrences, and there are four occurrences in the
Laytonville vicinity (Peterson et al. 1998). Four of
the seven localities have been discovered since 2001.
One historic occurrence (Round Valley) appears to be
extirpated, the other (Laytonville) was flourishing in
1998, leading Peterson et al. (1998) to state “... these
trees probably represent the optimum of currently
available habitat for the species”. Prior to 2001, the
abundant Laytonville locality was the only California
locality known to be extant.
The nine Lake Pillsbury occurrences have a
number of occupied trees, and abundance of Sulcaria
badia is moderate to high for a given tree (Toren
2004). In contrast, the Hawkins Creek and Campbell
Ridge localities have fewer than 10 thalli reported
each, and the Orleans occurrence is represented by a
single thallus. These, however, were detections made

from litterfall, and it should be borne in mind that
because of the nature of the forest (mid- to latemature Douglas-fir mixed with black oak, in high
canopy cover [>60%] stands), there may be a
significant additional presence in the canopy, that
was not visible from the ground. Even so, it is a
habitat that is wildly different from the Quercus
garryana community that was previously assumed to
be optimal, but it will be time-consuming and/or
expensive to discover how abundant Sulcaria badia
is in this habitat.
The two collections in ASU of Sulcaria badia
are from Fremont National Forest in Oregon and
Lassen National Forest in California. These
collections were reviewed by Peterson in 1996 and
determined to be Bryoria pseudofuscescens
(Schroeder 2006).
For all localities, there is a distinct possibility
that numerous thalli exist in the crowns of trees
above and adjacent to the understory locations where

Figure 2. Sulcaria badia in the Pacific Northwest.
Triangles represent historical sites that are
presumed extirpated; circles represent extant sites
or localities.
Sulcaria badia is found, since the surveys during
which the detections were made were all groundbased. It is also possible that this lichen is overlooked
and undercollected, since it shares certain
morphological and habitat characteristics with other
lichens, and can co-occur with or be dismissed as
Alectoria sarmentosa, Bryoria spp. and Usnea spp..

The modest distance (135 miles) between the
northern- and southernmost occurrences in California
adds credence to this possibility. But given the
distinctiveness of the deep spiraling pseudocyphellae
and the small number of ground detections made, it
seems reasonable to assume that occupied trees are
few, and the lichen is at least uncommon. Wineteer
(2004) states that survey intensity around the Snider
& Berry Creek BLM areas has been high, and only
two localities have been found.

47


BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 13 (2), 2006
campground building, etc. Conversion from oak
POPULATION TRENDS
woodland to residential usage has a twofold impact:
Peterson et al. (1998) reported thirteen total, but ten
removal of trees and the air quality issues resulting
extant occurrences of Sulcaria badia in the Pacific
from the increased presence of automobiles.
Northwest, with five extant and one possibly
Conversion to agricultural usage has
extirpated in California. The most
the same threats, with additional
current information for California,
impacts from the use of fertilizers and
assuming no new extirpations, is
herbicides.

that there are six general localities
(Figure 2), and nineteen discrete
PROTECTION
occurrences, using the ¼ mile rule
Status: Sensitive plant in Region
(CNPS 2006; this counts nine
5 of the US Forest Service
occurrences at Lake Pillsbury and
(California), 2006. Oregon Heritage
five at Laytonville). This increase
Program Rank G2 S2. Oregon Natural
should not be interpreted as a
Heritage Program List 1 (1 = contains
population trend, since it represents
taxa that are threatened with
new detections and probably not
extinction or presumed to be extinct
new recruitment. The lack of any
throughout their entire range), May
systematic
monitoring
efforts
2004 (elevated from List 2 in 2001).
makes an assessment of population
Washington
Natural
Heritage
trends impossible at this point.
Program List 1(Critically imperiled
Of the eleven extant localities

because of extreme rarity [5 or fewer
in Table 1, seven are composed of
occurrences, or very few remaining
more than one occupied tree, and 4
Figure 3. Sulcaria badia on Garry
individuals], or because of some
consist of fairly large populations
oak at Lake Pillsbury. Photo by
factor of its biology making it
(est. < 100 occupied trees) of
David Isle, USFS.
especially vulnerable to extinction) in
Sulcaria badia. This patchy2001, but the current Washington
clumpy distribution pattern at the
non-vascular
publication
is not available.
landscape level is highly typical of lichens with
The status of Forest Sensitive plant in the Forest
dispersal limitations, i.e. those that reproduce via
Service in Region 5 affords Sulcaria badia
large propagules. S. badia, as far as is known,
protections on Forest Service lands in California; the
disperses solely via thallus fragments.
individual Forests must manage known populations
to avoid a trend towards Federal listing with the Fish
THREATS
& Wildlife Service. This affects localities at Lake
History: Peterson et al. (1998) hypothesize that the
Pillsbury, Orleans, Campbell ridge and Hawkins

Round Valley occurrence may have become
Creek. However, the Orleans, Hawkins Creek and
extirpated because of marginal drier habitat coupled
Campbell Ridge localities are on matrix lands, and
with a decline in air quality caused by increased
are available for management, subject to mitigations
human presence (more fireplaces and wood smoke in
for species’ persistence.
the winter, when the lichen is metabolically active).
The Laytonville locality is on land of unknown
The Dungeness area has undergone a conversion to
ownership, but appears to be privately owned. The
agriculture, which has reduced the amount of forested
Round Valley occurrence is also on private property,
land, and air quality has declined there, too, as
although the possibility of extirpation may render the
evidenced by the numbers of nitrophilous lichens
land ownership moot. These occurrences are afforded
present. In general, conversion to agriculture or
no protections at present.
residential threaten this lichen, by removal of
substrate trees or the decline in air quality.
CONSERVATION SUMMARY
Future:
Removal of occupied or potentially
Sulcaria badia has a scattered distribution throughout
occupied trees will reduce populations of Sulcaria
its range, which is from central Oregon to northern
badia, and can occur through real estate
California. It is known from three habitats: Quercus

development, road building or maintenance,
garryana grasslands, mature Douglas-fir/black oak

48


Sulcaria badia Sponsorship
Table 1: Known localities of Sulcaria badia in the Pacific Northwest. UTM coordinates are Zone 10, NAD 27. 6R = Six
Rivers herbarium; other abbreviations for herbaria follow Index Herbariorum. Empty cells represent unknown or no data.
*
*
UTM N
Elevation Most recent
Herbarium
Presumed
State
County
Location
UTM E
(ft.)
observation
extant?
date
CA
Mendocino
Round Valley
477xxx 4404xxx
1420
1897
US

no
CA
Mendocino
Laytonville
457xxx 4395xxx
1700
1997
HSC
yes
CA
Humboldt
Orleans
455xxx 4570xxx
1580
2006
6R
yes
CA
Lake
Lake Pillsbury
505xxx 4359xxx
1820
2002
herb. Toren yes
CA
Trinity
Hawkins Creek
456xxx 4524xxx
1247
2005

6R
yes
CA
Trinity
Campbell Ridge
450xxx 4528xxx
1981
2004
6R
yes
CA
Lake
Bucknell Creek
498xxx 4354xxx
1960
2002
none
yes
OR
Benton
Philomath
330
1934?
OSC
no
OR
Benton
Corvallis
479xxx 4934xxx
230

1997
OSC
yes
OR
Benton
W of Philomath
470xxx 4931xxx
330
2003
OSC
yes
OR
Douglas
Reedsport
411xxx 4839xxx
131
1996
yes
OR
Jackson
Berry Creek
519xxx 4725xxx
1890
2003
OSC
yes
OR
Jackson
Snider Creek
505xxx 4711xxx

1650
2002
OSC
yes
WA
Clallam
Dungeness
490xxx 5332xxx
33
FH
no

forests, and shore pine/dune communities on the
immediate coast in Oregon. Population trends are
unknown, although two occurrences are presumed
extirpated, one of which causes a 225-mile
contraction in its range. Recent discoveries (post2002) on National Forest lands in northern California
have increased Sulcaria badia’s presence in
California, and it has recently become a Sensitive
plant on four National Forests in northern California.
These new occurrences are on matrix lands, and as
such are vulnerable to project-related disturbance, but
their status as Sensitive plants requires that
mitigations be written into planning documents.
SPECIFIC CONSERVATION RECOMMENDATIONS
Recommended Global Rarity Rank: G2G3
Species is narrowly distributed in the Northern
Hemisphere, restricted to the Pacific Northwest.
Localities adjacent to agricultural lands face loss of
vitality through declining air quality. Seven of eleven

localities consist of fewer than five occupied trees.
Recommended Global Threat Rank: .2
Possibly sensitive to air pollution, and some localities
are adjacent to agricultural lands, making these
vulnerable to human activities.
Recommended Local Rank (CA): S2S3.2
Small number of sites (six localities, nineteen
occurrences), only two localities have large number
of sites and/or occupied trees, only one of these
localities is on Federal lands.

Recommended Local Rank (OR): S2.2
Small number of sites (five localities, greater than
five sites), only two sites have large number of
occupied trees, both of these sites are on Federal
lands.
Recommended Local Rank (WA): SX
One historical occurrence in Washington state, which
is presumed extirpated.
Recommended CALS List (CA): 3
Sulcaria badia appears to be rare throughout its
range, and that range is narrow. Partially vulnerable,
since one large locality is on Federal land, but
abundances are typically low for a given occurrence.
Recommended CALS List (OR): 3
Same considerations as California, especially around
Medford BLM.
Recommended CALS List (WA): 1A
One historical site in Washington state, which is
presumed extirpated.

RELEVANT EXPERTS AND KNOWLEDGEABLE
LOCAL BOTANISTS
Eric Peterson, Ecologist/lichenologist
Nevada Natural Heritage Program
901 South Stewart Street, Suite 5002
Carson City, NV 89701

49


BULLETIN OF THE CALIFORNIA LICHEN SOCIETY 13 (2), 2006
Martin Hutten, Lichenologist
441 Hudson Road
Port Angeles, WA 98363
Bruce McCune, Professor of ecology and lichenology
Dept. Botany and Plant Pathology
Cordley 2082
Corvallis, OR 97331
STAKEHOLDERS FOR NOTIFICATION OF COMMENT
PERIOD
California Dept Fish and Game
Attn: Roxanne Bittman, CNDDB lead botanist
Wildlife & Habitat Analysis Branch
1807 13th Street Suite 2002
Sacramento, CA 95814
California Native Plant Society
Attn: Kristi Lazar, Rare Plant Botanist
2707 K Street, Suite 1
Sacramento, CA 95816-5113
Six Rivers National Forest:

Attn: Lisa Hoover, Forest Botanist
1330 Bayshore Way
Eureka, CA 95501
Klamath National Forest
Attn: Marla Knight, Forest Botanist
1312 Fairlane Road
Yreka, CA 96097-9549

LITERATURE (CITED OR OTHERWISE RELEVANT)
Brodo, I.M., D. Hawksworth. 1977. Alectoria and
allied genera in North America. Opera Botanica
42:1-164
Carlberg, T. 2005. Personal communication. Six
Rivers National Forest.
Hutten, M. 2004. Personal communication.

McCune, B. 2004. Personal communication.
Oregon Natural Heritage Program. 2004. Rare,
Threatened and Endangered Plants and Animals
of Oregon. Oregon Natural Heritage Program,
Portland, Oregon. 94 (98 pp. PDF)
Peterson, EB, DM Greene, B McCune, ET Peterson,
MA Hutten, P Weisberg, R Rosentreter. 1998.
Sulcaria badia, a rare lichen in western North
America. Bryologist 101(1): 112-115.
Toren, D., P. Nilles. 2003. Management
recommendations for Sulcaria badia Brodo & D.
Hawksw. a rare lichen occurring in the Lake
Pillsbury area of Lake County, CA; Mendocino
National Forest. Unpublished internal document,

USDA Forest Service, Mendocino National
Forest.
Wineteer, M. 2004. Personal communication.
Schroeder, R. 2006. Personal communication.
Toren, D. 2004. Personal communication. Mendocino
National Forest.

Mendocino National Forest
Attn: Lauren Johnson, Forest Botanist
825 N. Humboldt Ave.
Willows, CA 95988
Shasta-Trinity National Forest
Attn: Susan Erwin, Botanist
210 Main Street
Weaverville, CA 96093
Shasta-Trinity National Forest
Attn: Julie Nelson, Forest Botanist
3644 Avtech Parkway
Redding, CA 96002
Bureau of Land Management, Medford Field Office
Attn: Marcia Wineteer, Botanist
3040 Biddle Road
Medford, OR 97504

50

Sulcaria badia and Bryoria tortuosa drape branches
of an oak tree in roughly equal amounts, where S.
badia was rediscovered in the Laytonville area,
winter 1997. Photography by Eric Peterson.

Calicium adspersum,
Sponsorship for the CALS Conservation Committee
Eric B. Peterson
Nevada Natural Heritage Program
901 South Stewart Street, #5002
Carson City, NV 89701

EXECUTIVE SUMMARY
Calicium adspersum is a pin-lichen with a
yellowish pruina over the mazaedium, which
combined with the black spore mass gives the
head of the ascomata a dark-greenish cast. The
species has a large global range and is known in
North America from the Pacific Northwest in
areas of maritime-influenced cold climates. The
species appears to be quite infrequent through
its range within North America and at least some
parts of Europe. Only a single location is known
at present for the species in California, on state
lands with substantial management to retain the
old-growth forests currently at the site. Given
that there is only a single known site in
California and that appropriate habitats for this
sporadically occurring species are themselves
limited within California, the species qualifies for
ranking and listing by the California Lichen
Society and should meet criteria for protection
under the California Environmental Quality Act.

Recommendations are for rank G4.3 S1?.2 and
list 2. Suggested conservation is for assessment of the population at the known site and
management of forest and understory to provide
continuity of conditions around the site and that
care should be taken with prescribed fire to not
burn the trunks of trees inhabited by the species.

DESCRIPTION
Adapted from Tibell (1999): Lichenized calicioid.
Thallus grayish, generally verrucose. Ascomata a
stalked mazaedium, 0.8 – 1.4 mm tall, with a yellow
pruina over on the excipular rim and mixed with the
spores covering the mazaedium (sometimes faint)
and causing a greenish cast to the mazaedium.
Spores 13-17 um x 6-8 um, with a distinctive
ornamentation of spirally arranged ridges. Figure 1
(Figure C on back cover).
Note: Although the anatomy of Pacific
Northwest specimens matches well with European
specimens, the general morphology may be distinct
(Tibell pers. comm., email Dec. 2006), suggesting the
possibility of phylogenetic divergence, perhaps at a
subspecies level. Genetic study will be considered if
new specimens become available.
Similar species and distinguishing characteristics:
The genus is distinguished from other calicioid
genera most easily by the stalked ascomata with
black spores (under dissecting scope) and a

TAXONOMY

Accepted scientific name: Calicium adspersum
Pers.
Common name: none established; suggest “spiralspored guilded-head pin lichen”
Type specimen and location: Lectotype housed in
L, type location is Germany. (see Tibell 1999)
Synonyms: none known.

Figure 1 (repeated in color on back cover).
Calicium adspersum Pers. Photo from an Oregon
specimen, EBP #2737 (hb. Peterson). This
specimen shows a faint mixing of yellow pruina
into the mazaedium, resulting in a greenish
appearance. Scale bar = 1 mm. Photography by
E. B. Peterson.

51